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Transcript
Introduction
1
Oceanography, Geology, Biogeography, and Fisheries of the
Eastern Central Atlantic
1
2
1
3
by Merete Tandstad , Ana Maria Caramelo , Fabio Carocci , Kent Carpenter and Jonnell C Sanciangco
3
Introduction
his species identification guide focuses on the
marine species occurring in the eastern Atlantic
Ocean from Gibraltar in the north (36°00’N) to the
middle of Namibia in the south (23°00’S), and
westward to 30°00’W longitude (FAO Fishing
Area 34) and to 20°00’W longitude (FAO Fishing
Area 47) (Fig. 1). Estuarine species, as well as
species that only spend part of their life cycle in
marine waters (anadromous and catadromous
species) are also included.
T
The West African coast encompasses some of
the most productive areas in the world, supporting
important fishery resources and a unique diversity.
Fisheries constitute an important activity
throughout the region and contribute to
food security and economic revenue for
the people and countries from Morocco
in the north to Namibia in the south. The
fisheries are of a diverse nature and the
type of fishing craft used ranges from
small scale dug-out canoes, through
larger motorized canoes and coastal
fleets to large industrial vessels. Some
of the area covered by this guide
encompasses the greatest diversity of
marine organisms exploited by fisheries
in the eastern Atlantic Ocean.
case by case basis (FAO, 2009; Garcia et al.,
2010).
Industrial development in the coastal zone of
these areas, as well as migration of people from
inland rural areas to the coastal industrial centres,
has led to increasing pressure on the coastal
environment and habitat. Over the past 2 to 4
decades, marshes, swamps and mangroves have
been degraded and lost through natural factors
such as drought, but also through human activities
such as unsustainable agricultural practices,
urbanization, mining and other industries and
modification of rivers that has reduced water supply
The area covered is also diverse
culturally, politically, and geographically,
with many distinct marine regions and
several islands and island groups.
With many jurisdictional units it
becomes clear that the countries of the
region are faced with the challenge of
management of shared stocks. Many of
the main pelagic stocks and some of the
demersal stocks are distributed across
several EEZs. This has consequences
for the management of the fisheries
exploiting these resources and implies
that agreements (bilateral, subregional,
etc.) would need to be considered on a
Fig. 1 Area covered by this guide
1/
2/
3/
Marine and Inland Fisheries Branch, FAO Fisheries and Aquaculture Department, Rome, Italy.
Consultant, Marine and Inland Fisheries Branch, FAO Fisheries and Aquaculture Department, Rome, Italy.
Department of Biological Sciences & IUCN/Conservation International Global Marine Species Assessment, Old Dominion
University, Norfolk, Virginia, USA.
2
The Living Marine Resources of the Eastern Central Atlantic
to wetlands and marine areas (Heileman and
Tandstad, 2008). Overexploitation of natural
resources is also a fundamental problem in many
areas (FAO, 2008).
Purpose of this Guide
The purpose of this guide is to provide an
accurate tool to identify, to the appropriate taxonomic
level, those organisms that are of potential use or
likely to be captured by marine fisheries in the region.
The species included are those considered to be of
present or potential interest to fisheries, or those that
may be important in fisheries production or to
fisheries management in general.
It is hoped that this identification tool will benefit
fisheries workers gathering catch statistics and
resource assessment information, marine or fishery
biologists researching information pertinent to
resource management as well as the fisheries
administrations responsible for managing the
fisheries targeting these resources.
An additional objective of this guide is to provide a
tool that can be used to help documenting the
diversity of marine macrofauna in the region. This is
important also in relation to understanding the extent
of the biodiversity likely to be affected by fisheries.
Therefore, this guide is comprehensive and also
includes, to the extent possible, species that are not
commercially important.
This guide expands upon and updates the FAO
Species Identification Sheets for Fishery Purposes
for the Eastern Central Atlantic (Fischer et al., 1981).
All species accounts with distribution maps have
been revised and updated, and new species have
been added, expanding the coverage from 681 to
747 species. This guide is the geographic
complement to the FAO Identification Guide to the
Western Central Atlantic (Carpenter, 2002) and
together, the 2 guides provide coverage of
subtropical and tropical marine species across the
Central Atlantic Ocean.
Geology and Geomorphology
The western side of the African Plate dominates
the geology of the eastern Atlantic (Fig. 2).
Specifically this area includes the area bordered to
the north by the Azores-Gibraltar ridge, to the west
by the mid-Atlantic ridge and to the south by the
Walvis ridge and Cape basin.
The rupture of the Gondwana continent in
Mesozoic time and the gradual opening of the
Atlantic and Indian Oceans at the expense of the
Tethys Sea shaped the African plate. This was
followed by the Cenozoic opening of the Red Sea
seafloor spreading along the mid-Atlantic ridge.
This caused the separation of South America from
Africa, a process which lasted over a period of 100
Ma, from early Jurassic to middle Cretaceous
(Schwartz, 2005).
The principal feature of the bottom topography
of the Atlantic Ocean is the mid-Atlantic ridge, a
submarine mountain chain that extends from
Iceland in the north to approximately 58° south
latitude. The mid-Atlantic ridge separates the
Atlantic Ocean in two and transverse ridges running
between the continents and the mid-Atlantic ridge
divide the ocean floor into numerous basins.
Fig. 2 Major lithosphere plates of the world (source: Carpenter, 2002)
Introduction
3
To the east of the mid-Atlantic ridge
lie a number of large basins of relatively
even profile, interrupted by extended
ridge systems (Schwartz, 2005). To the
north of the eastern central Atlantic
region the larger basins include the Cape
Verde and Canaries basins whereas the
largest south Atlantic basins are the
Angola and Cape basins. The latter
basins are separated by a submarine
mountain chain, the Walvis ridge, which
runs from the continental shelf at latitude
20°S
(northern
Namibia)
in
a
southwesterly direction for more than
2 500 km towards the mid Atlantic ridge
(Fig. 3).
The basins are abyssal plains filled
by mud of organic calcareous origin and
red clay; and the ridges are rocky abyssal
hills. Large ancient volcanoes are found
in isolation or in rows and rise to form
seamounts and islands (Schwartz,
2005). This system of basins and ridges
influence the circulation of deeper waters
off the eastern central Atlantic.
Coastal Features and Shelf
Topography
The Atlantic coast of Africa may be
divided into three main zones: the coast
from Cape Spartel (Morocco) to Cape
Vert (Senegal) influenced by the cool
Canary Current; the humid tropical coast from Cape
Vert to beyond the Congo River, dominated by the
Guinea Current; and the coast from Angola to the
Cape of Good Hope characterized by the cool
Benguela Current.
Fig. 3 Bottom topographical features of the
eastern central Atlantic
One coastal feature of particular interest are the
mangroves that are found along the tropical parts of
the West African coast, from Mauritania in the north
to Angola in the south. Some 70% of all African
mangroves can be found in just five countries (FAO,
2007), including Nigeria, Guinea and Cameroon on
the Atlantic coast of Africa. Well developed
mangroves can be found in large river deltas, in
lagoons, along sheltered coastlines and on tidal flats
(FAO, 2007). These mangroves are important
nursery grounds for many fish and crustacean
species.
Another important characteristic, influenced
amongst others by the drainage of many major
African rivers into the eastern central Atlantic
(e.g. Cunene, Congo, Niger, Volta, Gambia,
Casamance), are the large estuarine areas and
coastal lagoons and the many wetlands that are of
considerable value within their extensive delta
systems. The lagoons in the Gulf of Guinea have
developed into a dominant feature of the coastline
and are important reservoirs for biological diversity
(Ajao et al., 2009).
The continental shelf along the coast of West
Africa although variable in width and depth is in
general narrow. The continental shelf in the CECAF
region (Area 34) is typically between 40 and 55 km
wide except in the area between latitudes 24°N and
20°N (Dakhla to Nouadhibou), and the area
between Dakar (15°N) and Freetown (8°N) where
the shelf is about 180 km wide (Brainerd, 1980). Off
southern Angola and Namibia the continental shelf
is narrow, being at its widest off the mouth of the
Orange River and off Walvis Bay, and narrowest off
the Cunene River to Cape Frio (Sakko, 1998 as
referred to in Palomares and Pauly, 2004; Bianchi
et al., 1999).
Most of the western coast of Africa is dominated
by terrigenous deposits, with the exception of a
small region off Ghana and Côte d’Ivoire (Longhurst
and Pauly , 1987).
4
The Living Marine Resources of the Eastern Central Atlantic
Oceanography
Winds, Currents and Upwellings
Wind intensity and direction significantly influence
the circulation of the surface water layers and the
oceanography of the West African region, and
subsequently also the productivity of the coastal
waters. These factors act at various times and space
scales ranging from region-wide seasonal changes to
local events of short duration.
The coastal environment along the west coast of
Africa is, however, not uniform, and this gives rise to
regional differences in productivity and biodiversity.
One unique feature of the West African coast is the
presence of 2 of the world’s 4 eastern boundary
current systems, the Canary current and the
Benguela current systems. Eastern ocean
boundaries are often referred to as classical coastal
upwelling regions (Bakun, 1996) and are
characterized by upwelling of cold nutrient-rich water
along the coast. These areas are important centres of
plankton production which again supports important
biodiversity and biomass of marine species
particularly pelagic fish (Longhurst and Pauly, 1987;
Bakun, 1996; Shannon and O’Toole, 1999).
The Canary Current branches south from the
North Atlantic Current and flows along the African
coast from north to south between 30°N and 10°N
and offshore to 20°W (Fedoseev, 1970). At the level
of Senegal the current moves away from the coast
and flows westwards joining the Equatorial Counter
Current coming from the south (Fig. 4). The
upwelling of cool, nutrient rich water along the
northwest African coast is the result of alongshore
wind forcing from the northeast trade winds, and
vary with season and latitude (Wooster et al., 1976).
In the Canary Current region major upwelling
occurs between 23°N and 25°N and occurs
year-round from Cap Blanc and northwards (FAO,
2009). South of Cap Blanc, upwelling is limited to
winter and spring due to the northward migration of
the Azores high during summer. The position and
strength of the Azores anticyclone governs the wind
and current system along the northwest African
coast and subsequently the intensity and area of
upwelling along the coast. The border between the
northeast and southeast trade winds, the
Intertropical Convergence Zone (ITCZ) is at its
southernmost limit in January–February (around
5°N) leading to an expansion of the up welling area
in this period. In the period June–August the ITCZ is
found further to the north and the limit of the
upwelling area retreats northwards.
The Benguela Current flows parallel to the
southwest African coast in a north to northwesterly
direction from approximately Cape Point in the
south until about 24°S where most of the current
bends northwest Streams of the Benguela current
proceed northward along the shore and its northern
limit corresponds to the frontal zone between the
Fig. 4 Major surface currents in a) February and b) July
(adapted from Wauthy, 1983 as shown in Schneider, 1990)
Introduction
5
Benguela and Angola current off
southern Angola (17°S to 13°S) (Figs
4 and 5). (Moroshkhin et al., 1970;
Wedepohl et al., 2000). The
prevailing winds are responsible for
strong Ekman transport and the
resulting coastal upwelling of cool,
nutrient-rich water that stimulates
primary productivity that sustains
large zooplankton production and
pelagic fish stocks.
The coastal upwelling area of the
Benguela current ecosystem extends
from the Cape in South Africa,
northwards along the coast of
Namibia and into southern Angola
(Shannon and O’Toole, 1999). The
intensity is not uniform in space and
time because of short term and
seasonal differences in the wind
regime and differences in coastline
and shelf topography (Bianchi et al.,
1999). Major centres of upwelling in
the Benguela include Cape Frio
(northern Namibia), off Luderitz in
southern Namibia and off the western
cape (South Africa), the first of which
is included in the area covered by this
guide.
Fig. 5 Major surface currents in the Benguela
The circulation patterns of the
waters of the Gulf of Guinea are
influenced by both the Canary and
1970), Kelvin waves (Picaut, 1983; Verstraete,
Benguela currents and also by the south Equatorial
1992), and cyclonic turbulent eddies (Marchal and
and Guinea Currents (Fig. 4). The latter flows
Picaut, 1977).
eastwards from Senegal to the Bight of Biafra and is
fed by the Equatorial Counter current and a branch of
The main upwelling region of the Gulf of Guinea
the Canary Current (Schneider, 1990). While the
extends from the coast of Côte d’Ivoire to Ghana.
northern border of the Guinea Current is distinct, but
The upwelling off the coasts of Ghana and Côte
with seasonal fluctuations, its southern boundary is
d’Ivoire is seasonal, with intense upwelling from July
less well-defined (Binet and Marchal, 1993). The
to September weakening from about January to
westward flowing South Equatorial current originates
March (Bakun, 1978; Roy, 1995). Another area of
between Cape Lopez (Gabon) and 18°S (Angola)
upwelling is located off Congo and Gabon where
and influences the waters off the Southern Gulf of
the raising of the thermocline, cooling of surface
Guinea.
water and increased primary production has been
interpreted as a sign of upwelling.
The Guinea Current is also characterized by
areas of upwelling and increased biological
Figure 6 illustrates regional differences and
productivity, although the processes leading to these
seasonal changes in sea surface temperatures
upwellings differs from those observed in the eastern
along the west African coast in 2009 showing higher
boundary systems and the Guinea Current is unusual
temperatures in the Gulf of Guinea area in January
among upwelling regions in that there seems to be no
and the relatively cooler waters in July.
correlation between sea surface temperature and
wind patterns on a seasonal time scale
Biogeography
(Longhurst,1962; Bakun, 1978). According to
Voituriez and Herbland (1982) seasonal upwelling in
The west coast of Africa encompasses
this region is not induced by local wind stress. It has
temperate, tropical and subtropical eastern Atlantic
been suggested that this upwelling is rather created
marine biogeographic regions. The different
by geostrophic adjustment of isotherms (Ingham,
physical and oceanographic features, along the
6
The Living Marine Resources of the Eastern Central Atlantic
Fig. 6 Mean sea surface temperatures (SST) for 2009 in a) January and b) July
[source: Images on Sea Surface Temperature obtained from site http://podaac.jpl.nasa.gov maintained by the
NASA JPL Physical Oceanography DAAC, Pasadena, CA (2010)].
coast influence the nature of the biota, leading to
regional differences in species dominance and
diversity along the coast. The West African
continental shelf represents the smallest tropical
shelf area of any of the four tropical regions. Northern
and southern oceanic gyres direct cool currents along
this coast, converge, and flow eastward to warm up
again on the path toward the Americas. This elevates
the depth of the thermocline along the coast of West
Africa, limiting the area of the shelf that tropical
organisms can thrive on and depresses marine
biodiversity compared to other tropical regions
(Briggs, 1974). Factors such as latitude and depth
also influence fish species diversity and species
richness generally increases as latitude decreases
and decreases with depth (Moyle and Cech, 2000).
Longhurst (1958) considers the absence of
reef-building corals in many tropical waters and
explains the predominance of sand, silt and shells in
bottom deposits. The very limited occurrence of reef
building corals in the eastern Atlantic can be
explained by conditions such as heavy rainfall and
numerous rivers carrying high volumes of fresh water
(thus lowering salinity) and silt into coastal waters.
These
conditions,
together
with
thermal
characteristics of the shelf waters elsewhere along
the west African coast combine to prevent the
formation of true coral reefs (Spalding et al., 2001).
Despite this, limited coral communities (mainly
restricted to the southern Gulf of Guinea) do exist
and the biota includes some typical coral reef
species that thrive on rocky reefs. The origins of this
reef fauna are primarily a result of dispersal in
relatively recent geological timescales, although
there are examples of Tethyan origins as well
(Floeter et al., 2008). Deep-water corals (e.g.
Lophelia spp.) are found in the deeper shelf and
slope area in patches throughout the region (Global
Distribution of Cold Water Corals. 2005.
UNEP-World Conservation Monitoring Centre.
http://data.unep-wcmc.org/datasets/1) and dead
coral banks are also occurring throughout the
region (Longhurst and Pauly, 1987).
Patterns of Diversity
Generalized patterns of biodiversity distribution
based on the species included in these volumes
were obtained by combining the range maps of
individual species. The reader must be aware that
the maps used are generalized maps that do not
take into consideration seasonal variations or
differences in distribution at different life stages nor
necessarily the actual observed depth ranges for
demersal species. The composite distribution of the
species included in this Guide shows that there are
three main concentrations of marine biodiversity
Introduction
7
along the west coast of Africa (Fig. 7).
The area of highest species diversity is
located in waters surrounding Senegal
and the area southeast of the Canary
Islands.
A peak in species richness off the
coast of Senegal that may be partially a
result of the proximity of the widest shelf
area on the west coast of Africa that
extends south from Senegal to Sierra
Leone. Another factor contributing to this
peak is also a number of warm temperate
species that extends only as far south as
Senegal (Briggs, 1974). However, as
mentioned previously, the maps used for
this analysis show generalized distribution
patterns that do not take into consideration
spatial and temporal changes based on
changes in environmental conditions.
Mixing of warm temperate and tropical
species is not detected in our analysis in
the southern part of the region because
our area of coverage of the identification
guide did not include the southern limit of
Africa. A second peak in diversity occurs
in the vicinity of the Canary Islands and
most likely represents a mixing of both
continental and island species that occur
in close proximity to the main coast of
Africa.
Fig. 7 Pattern of species richness in western Africa for
different taxonomic groups. All taxa (fishes, invertebrates,
and marine turtles)
industrial national fleets, the EEZs of the countries
in the area also accommodate distant water fleets
The southernmost concentration corresponds to
principally from Europe and Asia. Many of the
the equatorial waters of the Gulf of Guinea from
fisheries are typically multispecific thus posing
northern Angola northwards to Senegal. This is
additional challenges for assessment and
equatorial coastal biota with biodiversity attenuating
management.
in the offshore islands of São Tomé and Príncipe
(Fig. 7).
The total catches reported in FAO Fishing
Area 34 of the eastern central Atlantic was around
Diversity was found to be lower to the north and
4.1 million tonnes in 2012, below the 2008–2012
south of the study area corresponding to the more
average of around 4.2 million tonnes (Fig. 8). In the
temperate and subtropical waters of the northern
southern Atlantic (northern part of FAO Fishing
area of the Canary – and the Benguela Current
Area 47) catches were around 845 000 tonnes in
regions. Lower diversity in productive areas as
2012, above the 2008–2012 average of around
compared to oligotrophic ecosystems have been
739 000 tonnes and showing an increasing trend
noted by several authors (Margalef, 1974, 1997; Rex
since 2009 (Fig. 9).
et al., 2000 as referred to in MacPherson, 2002).
Latitudinal gradients in species richness have also
Around 300 species or groups of species were
been described, MacPherson (2002) noted that
reported
from FAO Fishing Area 34 during the
benthic taxa are strongly influenced by hydrographic
1975–2012 period by a total of 23 coastal states
events (especially upwelling processes and river
and 35 distant water fishing nations. In the northern
discharges) whereas pelagic species were less
part of FAO Fishing Area 47, 20 fishing nations
influenced by these coastal events.
(including the United Kingdom and South Africa) in
addition to Angola, Namibia and St Helena have
Fisheries
been reporting catches.
The eastern central Atlantic area is rich in
Table 1 shows the average landings for the
fisheries resources, including locally important
period 2008 to 2012 of the most important
resident stocks as well as transboundary, straddling
taxonomic units from FAO Fishing Areas 34
and migratory stocks. In addition to small scale and/or
8
The Living Marine Resources of the Eastern Central Atlantic
Fig. 8 Landings of main ISSCAAP4 groups in FAO Fishing Area 34
(source: FAO FishstatJ)
Fig. 9 Landings of main ISSCAAP groups in FAO Fishing Area 47 (northern part)
(source: FAO FishstatJ)
and 47. The small pelagics species notably the
clupeids sardine (Sardina pilchardus) and
sardinellas (Sardinella spp.) dominate the catches
in FAO Fishing Area 34, whereas in the northern
part of FAO Fishing Area 47 it is the Cape horse
mackerel (Trachurus capensis), the hakes
(Merluccius capensis and M. paradoxus), and the
clupeids that are the dominant species groups. The
fluctuations in total catches are thus highly sensitive
to changes in the catches of these species groups.
The composition of the fleet exploiting the main
resources has varied over time, and in FAO Fishing
Area 34 the coastal states have steadily developed
their national fisheries over the last decades
(Caramelo and Tandstad, 2005). Figure 10 illustrates
4/
the relative importance of landings of the different
fishing nations in the eastern central Atlantic, based
on the average landings by country for the period
2008 to 2012 as reported to FAO. The coastal
states account for around 66% of the catches in
FAO Fishing Area 34 whereas in the northern part
of FAO Fishing Area 47, Namibia and Angola
account for around 90%. The main coastal fishing
countries include Morocco, Senegal and Mauritania
to the north; Ghana, Sierra Leone and Nigeria in the
Gulf of Guinea and Namibia and Angola to the
south. Of the distant water fishing nations, Belize,
and the Russian Federation report the highest
landings in Area 34 whereas Spain, Taiwan
Province of China, Japan and the Russian
Federation represent the distant water fleets with
International Standard Statistical Classification of Aquatic Animals and Plants (ISSCAAP)
Introduction
9
the highest average catches in the
northern part of Area 47 during the
period 2008–2012 (Fig. 10).
Main Resources and State of
Stocks
Regional assessments of the state
of the main stocks in FAO Fishing Area
34 are traditionally carried out by the
pelagic and demersal Working Groups
of the Fisheries Committee for the
Eastern Central Atlantic (CECAF).
These Working Groups normally divide
Area 34 into two subareas, the northern
CECAF subarea from Morocco down to
the southern part of Senegal and the
southern subregion from Guinea-Bissau
to Angola, including Cape Verde and
São Tomé and Principe. The section
below reflects this subdivision. In
Southeast Africa national assessments
are carried out on a regular basis.
Regional assessments and scientific
surveys are also carried out through the
Benguela Current Commission.
Northern CECAF Area
Sardine (Sardina pilchardus) is the
Fig. 10 Countries landing in FAO Fishing Area 34 and
dominant species in the catches off
Northwest Africa5, constituting about FAO Fishing Area 47 (northern part). Average from 2008–2012
36% of overall main pelagic catch for
(Merluccius merluccius, M. senegalensis and
this subregion in 2011 followed by the sardinellas
M. polli) in 2012 represent on average 7% of total
(principally Sardinella aurita but also Sardinella
maderensis) that constituted 26%. Cunene horse catch of all demersal species analysed. The
deep-water
rose
shrimp
(Parapenaeus
mackerel (Trachurus trecae) is the most important
longirostris) and the southern pink shrimp
species of horse mackerels (FAO, 2013). Other
(Penaeus notialis) are also important in the region
important species include the Atlantic horse mackerel
and constitute 10% of catches of the demersal
(Trachurus trachurus), the chub mackerel
(Scomber
japonicus), anchovy (Engraulis species studied (FAO/CECAF, 2014b in
encrasicolus) and the bonga shad (Ethmalosa preparation).
fimbriata).
In general the small pelagic stocks in the
northern CECAF region have been the target of an
The most important group of demersal species
increasing demand for exploitation over the last
analysed by the FAO/CECAF Working Group on
decades. Recent assessments have shown that
demersal resources in the northern CECAF area is
stocks of Sardinella, in particular S. aurita as well
cephalopods, principally the common octopus
as the stocks of horse mackerel (Trachurus
(Octopus vulgaris). The common octopus is the
trecae) are overexploited in the subregion. Chub
focus of highly specialized fisheries on two stocks off
mackerel
(Scomber
japonicus), anchovy
Dakhla and Cape Blanc.
(Engraulis encrasicolus) and bonga shad
The red pandora (Pagellus bellottii), widely
(Ethmalosa fimbriata) are fully exploited. The
distributed in the West African zone, was the most
sardine to the south of Cape Boujador is not fully
important species in terms of catches of the demersal
exploited (FAO, 2011; FAO/CECAF, 2012).
fish (excluding hake) studied by the Working Group
Many of the commercially important demersal
until 2003. After this year, catfishes (Arius spp.) start
stocks in this region are in general intensely
to be as important as red pandora. Catches of hakes
5/
From Morocco to the South of Senegal
10
The Living Marine Resources of the Eastern Central Atlantic
exploited (Caramelo and Tandstad, 2005; Tandstad
and Caramelo, 2011). The assessments carried out
by the FAO/CECAF Working Group on demersal
resources off northwest Africa have shown that many
of the important demersal stocks such as the white
grouper (Epinephelus aeneus) stock in Mauritania,
the Gambia and Senegal is severely overexploited;
other overexploited stocks include the common
octopus (Octopus vulgaris) and shrimps
(Parapenaeus longirostris in Morocco and
Penaeus notialis in Senegal-Gambia) (FAO, 2011;
FAO/CECAF, 2012).
Southern CECAF Area
In the southern CECAF zone, between the northern
border of Guinea-Bissau and the southern border of
Angola, including the Cape Verde islands and the
islands of São Tomé and Príncipe, the most important
pelagic species in terms of landings are the two
sardinella species composed principally of the round
sardinella (S. aurita) but also including the Madeiran
sardinella (S. maderensis). Together they constituted
nearly 33% of the 2007 total landings of small pelagic
fish studied in the FAO/CECAF Working Group on the
Assessment of Small Pelagic Fish – Subgroup South in
October 2009. Anchovy (Engraulis encrasicolus),
bonga shad (Ethmalosa fimbriata) and horse
mackerels (principally Trachurus trecae) are also
important species landed in the southern CECAF
region (FAO/CECAF, 2013).
Of
the
demersal
species,
croakers
(Pseudotolithus spp.) from the sciaenid group made
up an average of about 37 000 tonnes in the period
2006–2010 with a contribution to the total catches of
the main demersal fish studied by the FAO/CECAF
Working Group on the Assessment of Demersal
Resources, Subgroup South, of about 14% in 2010.
Bigeye grunt (Brachydeuterus auritus) and
seabreams (Dentex spp.) are other important fish
species. Although low landings are recorded for
deep-water
rose
shrimp
(Parapenaeus
longirostris), southern pink shrimp (Penaeus
notialis) and cuttlefishes (Sepia spp.), these species
are considered highly valuable due to their high
commercial value.
Small pelagic resources are important but
fluctuating resources in this region where they are
mainly exploited by local small-scale or semiindustrial fleets, although some industrial fleets
operate in part of the subregion. They have mostly
been classified as non-fully to fully exploited, the
exception being the Sardinella aurita stock in the
western Gulf of Guinea (Côte d´Ivoire, Ghana, Togo
and Benin) and the southern stock of Cunene horse
mackerel (Trachurus trecae) (Gabon, Congo,
Congo DR and Angola) which were considered
overexploited (FAO, 2011; FAO, 2012; FAO/CECAF,
2013). The demersal resources in this region are
intensely exploited, and of the demersal species, 9
of the demersal stocks analysed by the last
FAO/CECAF Working Group were found to be
overexploited. This includes many demersal fish
stocks, such as West African croakers nei
(Pseudotolithus spp.) and others (FAO, 2011;
FAO/CECAF, 2012; FAO/CECAF, 2014a in
preparation).
Northern Part of FAO Fishing Area 47
In the northern part of FAO Fishing Area 47 the
large abundance of a few species characterises the
ichthyofauna of the pelagic zone in the southern
region.
The small pelagic fisheries of the region are
dominated by six taxonomic groups: cape horse
mackerel (Trachurus capensis) and cunene horse
mackerel (T. trecae), southern African pilchard
(Sardinops sagax, also still referred to as
S. ocellatus), southern African anchovy (Engraulis
capensis), sardinellas (round Sardinella aurita and
Madeiran or flat S. maderensis) and Whitehead's
round herring (Etrumeus whiteheadi) (Barros and
Cochrane, 2011).
Hakes (Merluccius capensis and M. paradoxus)
dominate the demersal fisheries. Other important
demersal species include seabreams (Dentex spp.)
and in particular the large-eye dentex (Dentex
macrophthalmus), shrimps and lobster (Penaeus
notialis, Parapenaeus longirostris and Panulirus
regius) and the West African geryon (Chaceon
maritae).
The pelagic species also dominate the landings
in this region, with the southern African pilchard
(Sardinops
sagax),
horse
mackerels
(Trachurus capensis and T. trecae), southern
African anchovy (Engraulis capensis) and
sardinellas (Sardinella aurita and S. maderensis)
contributing to the highest proportion of pelagic
landings, although catches of pilchard and horse
mackerels have decreased in recent years as
compared to earlier in this decade. Hake constitute
the demersal group with the highest reported
catches. Catches of hake have remained fairly
stable and have remained between 260 000 tonnes
and 320 000 tonnes since 1995 with a mean of
145 000 tonnes in the period 2008 to 2012.
Cochrane (2005) and Barros and Cochrane
(2011) noted that most of the commercially
important stocks within the region are classified as
being between fully and overexploited. Many (but
not all) of the currently overexploited stocks are
frequently the result of historical overexploitation
rather than current overfishing. A regional
assessment of the Cunene horse mackerel (T.
trecae) carried out through CECAF indicated that
this stock is overexploited.
Introduction
11
Fisheries dependent information, including accurate
information on catch or landings, is important for
fisheries management. However due to various
reasons including the difficulty in identifying
taxonomic units to species level, many countries
report species groups. As can be seen from Table 1,
this is also the case in the eastern central Atlantic.
This lack of taxonomic detail of catch data is a
challenge to fisheries scientists analysing the data for
management purposes, as species specific
parameters are frequently sought. The FAO CECAF
Assessment Working Groups and others have
pointed to the problems associated with low
resolution of taxonomic data and have stressed the
need to report catches to a lowest appropriate
taxonomic level for key species. This new addition of
Conclusion
the eastern central Atlantic guide is intended to help
The wide range of species and the high number of achieving this objective.
countries exploiting these resources in west Africa
pose a range of challenges for fisheries management.
The West African geryon stock from Angola and
Namibia (Chaceon maritae) is an important fishery
for the region. Assessment of this stock indicates that
it is currently overexploited, but probably recovering
slowly (MFMR, 2010). The shrimp fishery is also
important, in particular in Angola and includes both
deepwater (Parapenaeus longirostris and Aristeus
varidens) and shallow-water shrimps (Penaeus
kerathurus and other Penaeidae), the latter
important in the artisanal fishery. The Angolan stocks
of deep-water rose shrimp (Parapenaeus
longirostris) and striped red shrimp (Aristeus
varidens) are intensively exploited (Barros and
Cochrane, 2011).
Area 34
Statistical taxonomic unit
(common name)
Area 47 (north)
Average
landings
(04-08)
(tonnes)
Statistical taxonomic unit
(common name)
Average
landings
(04-08)
(tonnes)
Sardina pilchardus (sardine)
828 206
Trachurus capensis (Cape horse mackerel)
225 066
Trachurus spp. (horse mackerels)
418 083
Merluccius capensis, M. paradoxus (hakes)
152 183
Sardinella spp. (sardinellas)
355 840
Sardinella spp. (sardinellas)
74 025
Sardinella aurita (round sardinella)
281 392
Osteichthyes (bony fishes)
43 421
Scomber japonicus (chub mackerel)
237 325
Dentex spp. (seabreams)
32 665
Osteichthyes (bony fishes)
236 964
Sardinops sagax/S. ocellatus (southern African
pilchard)
22 233
Ethmalosa fimbriata (bonga)
235 548
Trachurus trecae (Cunene horse mackerel)
21 729
Sardinella maderensis (flat sardinella)
175 411
Pseudotolithus spp. (West African croakers nei)
19 692
Engraulis encrasicolus (anchovy)
170 287
Argyrosomus hololepidotus (southern meagre)
19 238
Katsuwonus pelamis (skipjack tuna)
161 109
Thunnus alalunga (albacore)
16 518
Thunnus albacares (yellowfin tuna)
88 414
Thunnus obesus (bigeye tuna)
13 972
Octopodidae (octopuses nei)
60 510
Ariidae (sea catfishes nei)
11 717
Ariidae (sea catfishes nei)
52 209
Ilisha africana (West African ilisha)
10 677
Thunnus obesus (bigeye tuna)
48 848
Scomber japonicus (chub mackerel)
10 554
Sepiidae, Sepiolidae (cuttlefishes and squids
nei)
31 959
Polynemidae (blue shark)
6 101
12
The Living Marine Resources of the Eastern Central Atlantic
Area 34
Statistical taxonomic unit
(common name)
Area 47 (north)
Average
landings
(04-08)
(tonnes)
Statistical taxonomic unit
(common name)
Average
landings
(04-08)
(tonnes)
Elasmobranchii (sharks, rays, skates nei)
30 743
Prionace glauca (blue shark)
6 043
Mugilidae (mullets nei)
28 684
Brachydeuterus auritus (bigeye grunt)
5 715
Brachydeuterus auritus (bigeye grunt)
27 469
Xiphias gladius (swordfish)
5 511
Sphyraena spp. (barracuda)
27 441
Lophius vomerinus (devil anglerfish)
4 972
Sciaenidae (croakers nei)
24 673
Genypterus capensis (kingklip)
4 403
Pseudotolithus spp. (croakers)
24 138
Pleuronectiformes (flatfishes nei)
3 538
Natantia (shrimps)
24 080
Thyrsites atun (snoek)
3 519
Polydactylus quadrifilis (giant African
threadfin)
23 085
Loligo reynaudi (Cape Hope squid)
3 466
Trichiurus lepturus (largehead hairtail)
22 743
Thunnus albacares (yellowfin tuna)
2 988
Pseudotolithus elongatus (bobo croaker)
21 081
Sciaenidae (croakers, drums nei)
2 699
Haemulidae (=Pomadasyidae) (grunt nei)
20 625
Merluccius capensis (shallow-water Cape hake)
2 232
Galeoides decadactylus (lesser African
threadfin)
18 559
Elasmobranchii (sharks, rays, skates nei)
2 216
Chloroscombrus chrysurus (Atlantic bumper)
17 567
Serranidae (groupers, seabasses nei)
1 567
Ilisha africana (African ilisha)
14 795
Austroglossus microlepis (West coast sole)
1 489
Cynoglossidae (tonguefishes nei)
14 777
Chaceon maritae (West African Geryon )
1 284
Rajiformes (skates and rays)
13 962
Pterogymnus laniarius (Panga seabream)
1 182
Sparidae (Porgies, seabreams nei)
13 645
Rajiformes (rays, stingrays, mantas nei)
1 148
Penaeus notialis (southern pink shrimp)
12 874
Pagrus spp. (Pargo breams nei)
1 002
Lutjanus spp. (snappers)
11 731
Zeus faber (John dory)
901
Nematopalaemon hastatus (estuarine prawn)
11 500
Loliginidae, Ommastrephidae (squids)
251
Euthynnus alletteratus (little tunny)
10 910
Engraulis capensis (southern African anchovy)
215
Pagellus bellottii (red pandora)
8 463
Parapenaeus longirostris (Deep-water rose
shrimp)
176
Merluccius merluccius (European hake)
7 575
Lepidopus caudatus (silver scabbardfish)
171
Pleuronectiformes (flatfishes nei)
7 262
Etrumeus whiteheadi (Whitehead's rund herring)
169
Octopus vulgaris (common octopus)
6 214
Jasus lalandii (Cape rock lobster)
116
Table 1. Landings (tonnes) of the most important statistical taxonomic units from FAO Fishing Areas 34
and 47 (North) from 2008 to 2012 ranked according to mean catch within the period.
Introduction
13
Map Data Sources
The projection used for the maps in this guide is a Cylindrical Equa-Area projection, with the central
meridian at 5°E, and the standardf parallel at 10°N. Country and continent borders are from the UN
Cartographic Unit, last updated in 2008. FAO Statistical Areas for Fishery Purposes were provided by
FAO Fisheries and Aquaculture Department. Bathymetric data are from the General Bathymetry Chart of
the Oceans (GEBCO) in grid format at resolution of 30 arc-seconds. Mean sea surface temperature maps
were obtained from site http://podaac.jpl.nasa.gov maintained by the NASA JPL Physical Oceanography
DAAC, Pasadena, CA in 2010.
Disclaimer
The designations employed and the presentation of material in the map(s) are for illustration only and do
not imply their expression of any opinion whatsoever on the part of FAO concerning the legal or
constitutional status of any country, territory or sea area, or concerning the delimitation of frontiers or
boundaries.
Literature
Ajao, E., Awosika, L., Diop, S., Koranteng, K., Soumare, A., & Snoussi, M. 2009. AoA Region: Western
African Seas 217–230 in UNEP/Dewa regional reports. www.unga-regular-process.org/images/
Documents/aoa%20annex%20iv%20regional%20summaries%30(3).pdf.
Bakun, A. 1978. Guinea Current Upwelling. Nature, 271, 147–150.
Bakun, A. 1996. Patterns in the Ocean. Ocean processes and marine population dynamics. California Sea
Grant College System. National Oceanic andd Atmopospheric Administration in cooperation with
Centro de Investigaciones Biológicas del Noroeste, La Paz, BCS México.
Barros, P. & Cochrane, K. 2011. Southeast Atlantic. In FAO. Review of the state of world marine fishery
resources, pp. 107–119. FAO Fisheries Technical Paper. No. 569. Rome, FAO. 334 pp.
Bianchi, G. 1993. Demersal assemblages of tropical continental shelves. A study based on the data collected
through the surveys of the R/V Dr. Fridtjof Nansen. Thesis for Dr. Scient. degree, University of
Bergen, Norway.
Bianchi, G., Carpenter, K.E., Roux, J.-P., Molloy, F.J., Boyer, D. & Boyer, H.J. 1999. FAO Species
identification field guide for fishery purposes. The living marine resources of Namibia. Rome, FAO.
1999. 265 pp.
Bianchi, G., Lundsør, E. & Hamukuaya, H. 2004. On Namibia’s marine fish diversity. In U.R. Sumaila,
D. Boyer, M.D. Skogen & S.I. Steinshamn, eds. Namibia’s fisheries: Ecological, Economic and
Social Aspects. Eburon Academic Publishers, The Netherlands. 363 pp.
Binet, D. & Marchal, E. 1993. The large marine ecosystem of shelf areas in the Gulf of Guinea: long-term
variability induced by climatic changes. In K. Sherman, L. Alexander & B. Gold, eds. Large marine
ecosystem V: Stress, mitigation and sustainability of large marine ecosystems, pp. 104–118.
Washington: American Association for the Advancement of Science.
Brainerd, T.R. 1980. Appendix 4. The demersal fisheries of Sierra Leone with notes on recent research
surveys. In FAO 1980: Report of the second ad hoc working group on coastal demersal stocks from
southern Mauritania to Liberia (CECAF statistical divisions 34.3.1 and 34.3.3) CECAF/ECAF
SERIES 80/23 (also available in www.fao.org/docrep/003/N9567E/N9567E00.htm).
Briggs, J.C. 1974. Marine Zoogeography. McGraw-Hill, Inc., New York.
14
The Living Marine Resources of the Eastern Central Atlantic
Caramelo, A.M. & Tandstad, M. 2005 Eastern Central Atlantic, In FAO. Review of the state of world marine
fishery resources, pp. 67–75. FAO Fisheries Technical Paper. No. 457. Rome, FAO. 235 pp.
Carpenter, K.E., ed. 2002. The living marine resources of the western central Atlantic. Volume 1:
Introduction, molluscs, crustaceans, hagfishes, sharks, batoid fishes, and chimaeras. FAO Species
identification guide for fishery purposes and American Society of Ichthyologists and Herpetologists
special publication No. 5. Rome, FAO. 2002. pp. 1–600.
Cochrane, K. 2005. Southeast Atlantic. In FAO. 2005. Review of the state of world marine fishery resources.
pp. 76–86. FAO Fisheries Technical Paper. No. 457. Rome, FAO. 235 pp.
FAO. 1990–2014. FAO Major Fishing Areas. ATLANTIC, SOUTHEAST (Major Fishing Area 47). CWP Data
Collection. In: FAO Fisheries and Aquaculture Department [online]. Rome. Updated 24 September
2008. [Cited 28 July 2014]. www.fao.org/fishery/area/Area47/en.
FAO. 1990–2014. FAO Major Fishing Areas. ATLANTIC, EASTERN CENTRAL (Major Fishing Area 34).
CWP Data Collection. In: FAO Fisheries and Aquaculture Department [online]. Rome. Updated 1
October 2004 [Cited 28 July 2014]. http://www.fao.org/fishery/area/Area34/en.
FAO. 2007. The world's mangroves 1980–2005. FAO Forestry Paper 153. FAO. Rome: 77 pp.
FAO. 2008. Fishery Committee for the Eastern Central Atlantic/Comité des pêches pour l’Atlantique
Centre-Est. Report of the fifth session of the Scientific Sub-Committee. Casablanca, Morocco, 4–6
December 2007. Rapport de la cinquième session du Sous-Comité scientifique. Casablanca,
Maroc, 4–6 décembre 2007. FAO Fisheries Report/FAO Rapport sur les pêches. No.869. Accra,
FAO. 2008. 50 pp. (also available in www.fao.org/docrep/015/i0225b/i0225b00.htm).
FAO. 2009. Protection of the Canary Current large marine ecosystem (CCLME). Project document
GCP/INT/023/GFF.
FAO. 2011. Fishery Committee for the Eastern Central Atlantic/Comité des pêches pour l'Atlantique
Centre-Est. Report of the sixth session of the Scientific Sub-Comité. Accra, Ghana, 7–9 September
2011. Rapport de la sixième session du Sous-Comité scientifique. Accra, Ghana, 7–9 septembre
2011. FAO Fisheries and Aquaculture Report/FAO Rapport sur les pêches et l'aquaculture. No. 985.
Accra, FAO. 2011. 79 pp.
FAO. 2013. Report of the FAO Working Group on the assessment of small pelagic fish off Northwest Africa.
Dakar, Senegal 21–25 May 2012/Rapport du Groupe de travail de la FAO sur l’évaluation des petits
pélagiques au large de l’Afrique nord-occidentale. Dakar, Sénégal 21–25 mai 2012. FAO Fisheries
and Aquaculture Report/FAO Rapport sur les pêches et l’aquaculture. No. 1036. Rome, FAO.
245 pp.
FAO. 2011–2014. Fisheries and aquaculture software. FishStatJ – software for fishery statistical times series.
(Capture Production 1950–2012). In: FAO Fisheries and Aquaculture Department [online]. Rome.
Updated 22 July 2014. [Cited 28 July 2014]. www.fao.org/fishery/statistics/software/fishstatj/en
FAO/CECAF. 2012. FAO Fishery Committee for the Eastern Central Atlantic/Comité des pêches pour
l’Atlantique Centre-Est. Report of the FAO/CECAF Working Group on the assessment of demersal
resources – Subgroup South/Rapport du Groupe de travail FAO/COPACE sur l’évaluation des
ressources démersales – Sous-groupe sud. Freetown, Sierra Leone, 9–18 October 2008.
CECAF/ECAF Series/COPACE/PACE Séries. No. 11/73. Rome, FAO. 311 pp.
FAO/COPACE. 2013. FAO Fishery Committee for the Eastern Central Atlantic/Comité des pêches pour
l’Atlantique Centre-Est. Report of the FAO/CECAF Working Group on the assessment of small pelagic
fish – Subgroup South. Accra, Ghana, 19–28 October 2009/Rapport du Groupe de travail
FAO/COPACE sur l’évaluation des petits poissons pélagiques – Sous-Groupe Sud. Accra, Ghana,
19–28 octobre 2009. CECAF/ECAF Series/COPACE/PACE Séries. No. 12/74. Rome, FAO. 172 pp.
Introduction
15
FAO/COPACE. 2014a. (in preparation). FAO Fishery Committee for the Eastern Central Atlantic/Comité des
pêches pour l’Atlantique Centre-Est. Report of the FAO/CECAF Working Group on the assessment
of demersal resources – Subgroup South/Rapport du Groupe de travail FAO/COPACE sur
l’évaluation des ressources démersales – Sous-groupe sud. Accra, Ghana, 15–24 November 2011.
CECAF/ECAF Series/COPACE/PACE Séries.
FAO/COPACE. 2014b. (in preparation). FAO Fishery Committee for the Eastern Central Atlantic/Comité des
pêches pour l’Atlantique Centre-Est. Report of the FAO/CECAF Working Group on the assessment
of demersal resources – Subgroup North/Rapport du Groupe de travail FAO/COPACE sur
l’évaluation des ressources démersales – Sous-groupe nord. Fuengirola, Spain, 18–27 November
2013. CECAF/ECAF Series/COPACE/PACE Séries.
Fedoseev, A. 1970. Geostrophic circulation of surface waters on the shelf of north-west Africa. Rapp. P.-V.
Reun. Cons. Int. Explor. Mer., 159, 32–37.
Fischer, W., Bianchi, G. & Scott, W.B., eds. 1981. FAO Species identification guide for fishery purposes.
Eastern Central Atlantic (Fishing Areas 34 and 47; in part). Canada Funds-in-Trust. Ottawa,
Department of Fisheries and Oceans, Canada, by arrangement with the Food and Agriculture
organization of the United Nations. Vol. 1–7, pag. var.
Floeter, S.R., Rocha, L.A., Robertson, D.R., Joyeux, J.C., Smith-Vaniz, W.F., Wirtz, P., Edwards, A.J.,
Barreiros, J.P., Ferreira, C.E.L., Gasparini, J.L., Brito, A., Falcón, J.M., Bowen, B.W., &
Bernardi, G. 2008. Atlantic reef fish biogeography and evolution. Journal of Biogeography, 35,
22-47.
Garcia, S., Tandstad, M. & Caramelo, A.M., eds. 2010. Symposium science and the challenge of managing
small pelagic fisheries on shared stocks in Northwest Africa, 11–14 March 2008, Casablanca,
Morocco/Symposium Science et défi de l’aménagement des pêcheries de petits pélagiques sur les
stocks partagés en Afrique nord-occidentale, 11–14 mars 2008, Casablanca, Maroc. FAO Fisheries
and Aquaculture Proceedings/FAO Comptes rendus des pêches et de l’aquaculture. No. 18. Rome,
FAO. 2010.
Ingham, M.C. 1970. Coastal upwelling in the northwestern gulf of Guinea. Bulletin of Marine Science, 20:
1–34.
Heileman, S. & Tandstad, M. 2008. Canary Current LME In K. Sherman & G, Hempel, eds. The UNEP large
marine ecosystems report: A perspective on changing conditions in LMEs of the world’s regional
seas. Nairobi, Kenya: UNEP. www.Ime.noaa.gov/LMEWeb/LME_Report/Ime_27.pdf. pp. 131–141.
Hempel, G., O’Toole, M. & Sweijd, N., eds. 2008. Benguela. Current of plenty. A history of international
cooperation in marine science and ecosystem management. Benguela Current Commission.
143 pp.
Longhurst, A. 1958. An ecological survey of the West African marine benthos. Fish. Publ. Colon. Off., Lond.,
(11):101 pp.
Longhurst, A.R. 1962. A review of the oceanography of the Gulf of Guinea. Bull. Inst. Afr. Noire, 24: 633-663.
Longhurst, A.R. & Pauly, D. 1987. Ecology of tropical oceans. Academic Press Inc., London, 389 pp.
MacPherson, E. 2002. Large scale species richness gradients in the Atlantic Ocean. Proc. R. Soc. Lond. B
2002, 269: 1715–1720.
Marchal, E. & Picaut, J. 1977. Répartition et abondance évaluées par écho-intégration des poissons du
plateau ivoiro-ghaneen en relation avec les upwellings locaux. J. Rech. Oceanogr., 2: 39–57.
Margalef, R. 1974. Ecologoía. Barcelona: Editorial Omega.
16
The Living Marine Resources of the Eastern Central Atlantic
Margalef, R. 1997. Our biosphere. Excellence in Ecology 10. Oldendorf, Germany: Ecology Institute.
Ministry of Fisheries and Marine Resources of Namibia (MFMR). 2010. An overview Namibia's
commercial Fisheries in 2009. Swakopmund, Namibia. 3 pp.
Monteiro, P.M.S., Spolander, B., Brundrit, G.B. & Nelson,G. 1998. Shellfish mariculture in the Benguela
system: estimates of nitrogen-driven new production in Saldanha Bay using two physical models.
Journal of Shellfish Research. 17(1): 3–13.
Moroshkhin, K. V., Bubnov, V.A. & Bulatov. R.P. 1970. Water circulation in the Eastern South Atlantic
Ocean. Oceanology, 10(1): 27–34.
Moyle, P.B. & Cech, Jr., J.J. 2000. Fishes. An introduction to ichthyology. Fourth edition. Prentice Hall, 612 pp.
Palomares, M.L. & Pauly, D., eds. 2004. West African marine ecosystems: Models and fisheries impacts.
Fisheries Centre Research Report, 12(7): 221 pp.
Picaut, J. 1983. Propagation of the seasonal upwelling in the eastern equatorial Atlantic. Journal of Physical
Oceanography, 13: 18–37.
Rex, M.A., Stuart, C.T. & Coyne, G. 2000. Latitudinal gradients of species richness in the deep-sea benthos
of the North Atlantic. Proc. Natl Acad. Sci. USA, 97: 4082–4085.
Roy, C. 1995. The Côte d’Ivoire and Ghana coastal upwellings: Dynamics and changes. In F. Bard & K.A.
Koranteng, eds. Dynamique et usage des ren sardinelles de l’upwelling cotier du Ghana et de la
Côte d’Ivoire. ORSTOM edition, Paris, France.
Sakko, A.L. 1998. The influence of the Benguela upwelling system on Namibia’s marine biodiversity.
Biodiversity and Conservation, 7: 419–433.
Schneider, W. 1990. FAO Species identification sheets for fishery purposes. Field guide to the commercial
marine resources of the Gulf of Guinea. Rome, FAO.1990. 268 pp.
Shannon, L.V. & O’Toole, M. 1999. Integrated overview of the oceanography and environmental variability
of the Benguela current region. BCLME Thematic report No. 2, UNDP, Windhoek. 57 pp.
Spalding, M.D., Ravilious, C. & Green, E.P. 2001. World atlas of coral reefs. University of California Press,
Berkeley.
Schwartz, M.L., ed. 2005. Encyclopedia of coastal science. Springer. 1211 pp.
Tandstad, M. & Caramelo, A.M. 2011. Eastern Central Atlantic. In FAO. Review of the state of world marine
fishery resources, pp. 67–75. FAO Fisheries Technical Paper. No. 569. Rome, FAO. 334 pp.
Verstraete, J.M. 1992. The seasonal upwellings in the Gulf of Guinea. Progress in Oceanography, 29: 1-60.
Voituriez, B. & Herbland, A. 1982. Comparaison des systèmes productifs de l’Atlantique tropical est: dômes
thermiques, upwellings côtiers et upwelling équatorial". Rapp. P.-v. Reun. Cons. int. Explor. Mer.,
180: 114–130.
Wauthy, B. 1983. Introduction à la climatologie du Golf de Guinée. Océanogr. Trop., 18(2): 103–138.
Wedepohl, P.M., Lutjeharms, J.R.E. & Meeuwis, J.M. 2000. Surface drift in the south-east Atlantic Ocean.
South African Journal of Marine Science, 22: 71–79.
Wooster, W.S., Bakun, A. & McLain, D.R. 1976. The seasonal upwelling cycle along the eastern boundary
of the North Atlantic. Journal of Marine Research, 34: 131–140.